The Metathoracic Spiracles in Some Ants and Wasps (Hymenoptera: Formicidae; Vespidae)
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Russian Entomol. J. 26(1): 49–62 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2017 The metathoracic spiracles in some ants and wasps (Hymenoptera: Formicidae; Vespidae) Ìåòàòîðàêàëüíûå äûõàëüöà íåêîòîðûõ ìóðàâüåâ è îñ (Hymenoptera: Formicidae; Vespidae) Elena B. Fedoseeva Åëåíà Á. Ôåäîñååâà Zoological Museum of Lomonosov Moscow State University, Bolshaya Nikitskaya Street 2, 125009 Moscow, Russia. Зоологический музей МГУ им М.В. Ломоносова, Большая Никитская ул., 2, 125009 Москва, Россия. KEYWORDS: Aculeata, Comparative anatomy, mesosoma, mesopleuron, metapleuron, spiracular mechanism. КЛЮЧЕВЫЕ СЛОВА: Aculeata, сравнительная морфология, мезосома, мезоплеврон, метаплеврон, механизм дыхальца. ABSTRACT. The second thoracic spiracle and the мы со скрытыми дыхальцами из обоих семейств adjacent skeleton were studied in twenty-two ant species Aculeata имеют запирающую дыхальцевую мышцу в from nine Formicidae subfamilies (Formicinae, Doli- дополнение к супрамезоплевральному склериту, choderinae, Myrmeciinae, Paraponerinae, Ponerinae, Ec- скрывающему истинное дыхальце. Предполагается tatomminae, Myrmicinae, Dorylinae and Ecitoninae) and гомология между повторно обнаруженной дыхаль- two wasp species from Vespidae (Vespa crabro L., 1758 цевой мышцей Aculeata и соответствующей мышцей and Dolichovespula sp.). Dissections followed by scan- метаторакального дыхальца Symphyta. Сравнение ning electron microscopy or photography revealed that крылатых Formicidae и рабочих муравьев со скрыты- the studied alate forms with concealed spiracles from ми дыхальцами продемонстрировало сходство меж- both Aculeata families possess the spiracular occlusor ду ними как по положению пластинок, покрываю- muscle in addition to the supramesopleural sclerite con- щих дыхальца (т.е. супрамезоплеврального склерита cealing the true spiracle. There is assumed to be a homol- и дыхальцевой лопасти, соответственно), так и по ogous relationship between the re-discovered spiracular положению дыхальцевых мышц. Поэтому дыхальце- occlusor muscle of Aculeata and the corresponding mus- вую лопасть рабочих муравьёв Formicinae, Dolicho- cle of the metathoracic spiracle in Symphyta. The com- derinae, Myrmeciinae, Paraponerinae, Ponerinae, Ecta- parison of winged Formicidae and ant workers with tomminae следует считать дериватом супрамезоплев- concealed spiracles demonstrated the similarities be- рального склерита, присущего крылатым самкам. В tween them both on the position of plates covering the свою очередь, открытый тип дыхалец у рабочих му- metathoracic spiracles (i.e. supramesopleural sclerite and равьёв Myrmicinae, Dorylinae и Ecitoninae согласует- spiracular lobe, respectively), and on the position of the ся с отсутствием супрамезоплеврального склерита у spiracular occlusor muscles. Accordingly, the spiracular крылатых форм из тех же подсемейств. lobe of worker ants of Formicinae, Dolichoderinae, Myr- meciinae, Paraponerinae, Ponerinae and Ectatomminae Introduction was determined to be a derivative of the supramesopleu- ral sclerite that is essential to their winged females. At the same time the exposed type of spiracles observed in As the metathoracic spiracles are inconspicuous or studied Myrmicinae, Dorylinae and Ecitoninae ant work- concealed, it is difficult to investigate their morphology ers corresponds with the absence of the supramesopleural and phylogeny in aculeate hymenoptera. Nevertheless, sclerite in the alate forms. their position is one of the important landmarks on the hymenoptera thorax [Reid, 1941]. In winged forms, the РЕЗЮМЕ. Второе грудное дыхальце и примыка- second thoracic spiracles are concealed beneath the bases ющий к нему скелет были изучены у 22 видов мура- of the hind wings; in wingless forms, the original position вьёв из 9 подсемейств Formicidae (Formicinae, of the spiracle is in the membrane between the meso- and Dolichoderinae, Myrmeciinae, Paraponerinae, Ponerinae, metapleuron. According to Reid, in many forms the Ectatomminae, Myrmicinae, Dorylinae и Ecitoninae), а second spiracle is at the dorsal termination of the me- также у двух видов ос из Vespidae (Vespa crabro L. sometapleural suture, and with very few exceptions it is 1758 and Dolichovespula sp.). Препаровка с последу- always at the junction of the dorsum and pleura. ющей сканирующей электронной микроскопией либо Meanwhile et Duncan [1939] noted that second thorac- фотосъёмкой показали, что изученные крылатые фор- ic spiracle of vespine wasps is covered by a small sclerite. 50 E.B. Fedoseeva He referred to it as the spiracular peritreme, but subsequent spiracle, were analysed in the course of compiling the comparison with other Hymenoptera suggested that it is a attributes of species, genera and subfamilies [Yoshimura, real sclerite [Tonapi, 1958]. Primarily it was attributed to Fisher, 2007, 2012; Boudinot, 2015]. Information about the posterodorsal corner of the mesopleuron [Duncan, the status of this trait in females is published infrequently 1939; Tonapi, 1958]. However, currently, based on data of [Ogata 1987, 1991]. However the internal structure of the symphytans, modern authors referred to sclerite as certain metathoracic spiracle in Formicidae species is not men- intersegmentalia that were affiliated with the mesopleuron tioned in modern publications. [Vilhelmsen, 2000; Vilhelmsen et al., 2010]. Besides vespine wasps, the sclerite that conceals or partly covers Materials and methods the second thoracic spiracle was detected in some other taxa of Aculeata. As a result, it acquired a variety of denominations: collar [Tonapi, 1958]; epimeral lobe [Ri- The examined material included two wasp species chards, 1977; Yoshimura, Fisher, 2007]; basalar lobe from Vespidae (Vespinae), Vespa crabro Linnaeus, [Bohart, Menke, 1976; Deyrup, Cover, 2004; MacGown 1758 and Dolichovespula sp., and twenty-three ant et al., 2014], supramesopleural sclerite [Vilhelmsen et species from nine subfamilies of Formicidae: Formici- al., 2010], spiracular sclerite [Boudinot, 2015]. nae, Dolichoderinae, Myrmeciinae, Paraponerinae, Pon- The mechanism of the second thoracic spiracle in erinae, Ectatomminae, Myrmicinae, Dorylinae and Aculeata representatives remains uncertain although many Ecitoninae (Table). Detailed information about speci- researchers have studied it. Most agree with certain sim- mens is presented in Appendix 1. ilarities between the second spiracle and the first one, in Table. List of examined Formicidae species which the aperture is regulated by a single occlusor Таблица. Список изученных видов Formicidae muscle. The principal disagreement concerns the exist- ence of the corresponding muscle. For example, there is Subfamily Species an occlusor muscle of the second thoracic spiracle in Dolichoderinae Azteca sp. several ant species [Nasonov, 1889; Tonapi, 1958, 1959; Dolichoderus attelaboides (Fabricius, 1775) Delye, 1965]. Duncan [1939], Tonapi [1958] and Rich- Dorylinae Dorylus orientalis Westwood, 1835 ards [1977] noted the presence of the occlusor muscle in Ecitoninae Eciton burchellii (Westwood, 1842) vespine wasps while Snodgrass [1925] declared its ab- Ectatomminae Ectatomma tuberculatum (Olivier, 1792) sence in honeybees. In addition to above mentioned Formicinae Ñamponotus (Tanaemyrmex) sp. aculeates Tonapi [1958] studied spiracular structures in Ñamponotus herculeanus (Linnaeus, 1758) Symphyta species. Most of his conclusions relating to the Formica aquilonia Yarrow, 1955 Formica exsecta Nylander, 1846 presence of an occlusor muscle of the second thoracic Formica lugubris Zetterstedt, 1838 spiracle in symphytans were corroborated [Vilhelmsen, Lasius niger (Linnaeus, 1758) 2000], but the muscle’s existence in aculeates requires Myrmeciinae Myrmecia sp. additional studies. The main problem is that it is such a Myrmecia piriformis Smith, F., 1858 minute muscle and is therefore difficult to detect. More- Myrmicinae Aphaenogaster gibbosa (Latreille, 1798) over, sometimes it “is not possible to discern between Atta sexdens (Linnaeus, 1758) concealment and absence of the spiracle because this Manica rubida (Latreille, 1802) fragile structure was often destroyed during dissection” Messor denticulatus Santschi, 1927 [Vilhelmsen et al., 2010: 72]. Myrmica lobicornis Nylander, 1846 Since the attributes of the metathoracic spiracle as Myrmica ruginodis Nylander, 1846 Pogonomyrmex badius (Latreille, 1802) well as the state of its associated sclerite are important for Paraponerinae Paraponera clavata (Fabricius, 1775) interpretation of Aculeata phylogeny, it was further eval- Ponerinae Diacamma geometricum Smith F., 1857 uated in this study. It was necessary to demonstrate the Ponerinae Dinoponera gigantean (Perty, 1833) adjacent skeleton and the components of the spiracle at least in representatives of the two families Vespidae and Formicidae. The number of species in Formicidae was Most of the material for the present study was pre- greater for the following reasons. Most Formicidae spe- served in 70% ethanol. Dried material was used in some cies have winged sex individuals (males and females) and instances. The dissections were carried out with tools for wingless workers (non-reproductive females), that mark- ophthalmic microsurgery (spring scissors, blade holder edly differed in thorax morphology. Moreover the thorax and forceps), pieces of razor blades and hair, and entomo- structure varies between Formicidae subfamilies. There- logical pins. Prior to dissecting the mesosoma, the head, fore it was necessary to study the representatives of legs, wings and metasoma were removed. Dissection